JournaloftheLepidopterists'Society 45(2), 1991, 135-141 A REVIEW OF APODEMIA HEPBURN I (LYCAENIDAE: RIODININAE) WITH A DESCRIPTION OF A NEW SUBSPECIES George T. Austin Nevada State Museum and Historical Society, 700 Twin Lakes Drive, Las Vegas, Nevada 89107 ABSTRACT. Apodemia hepburni Godman and Salvin (Lycaenidae: Riodininae) is reviewed. Anewsubspecies, Apodemia hepburniremota Austin, isdescribedfromsouth- ern Baja California, Mexico, based on 112 specimens. Both taxa of the species exhibit biphenism. Additionalkeywords: BajaCaliforniaSur, Sonora, Mexico,A. palmerii,A. murphyi. Apodemia hepburni is a small metalmark (Lycaenidae: Riodininae) described from Chihuahua, Mexico (Godman & Salvin 1886). In con- junction with a study of the phenotypically similar Apodemia palmerii (W. H. Edwards) (Austin 1987), I had the opportunity to examine series of A. hepburni in several major museums in the United States and those in a number of private collections. The degree of seasonal and geo- graphical variation I noted prompted this review. Throughout, butterfly size (mean and range in mm) is the length of the forewing from base to apex. Specimens indicated by "M" and "F" are male and female, respectively. Capitalized color names are after Smithe (1975, 1981). Apodemia hepburni hepburni Godman and Salvin (Figs. 1-4) Apodemia hepburni GodmanandSalvin 1886:468 (typelocality: PinosAltos, Chihuahua, Mexico); holotype male at The Natural History Museum, London (Miller & Brown 1981). Thespecieswasfigured andverybrieflydescribedfrom asingle malebyGodmanand Salvin (1886). I examined photographs of the type, a somewhat worn male without antennae and with a phenotype of the first brood. This specimen has six labels: a white hand-lettered label "HOLOTYPE/Apodemia/hepburni/Godman & Salvin/det. P. Ack- ery, 1989"; printed white labels with "B.C.A.Lep.Rhop./Apodemia/hepburni,/G. & S./ Godman-Salvin/Coll. 1914.-5.", "Pinos Altos,/Chihuahua,/Mexico/Buchan-Hepburn.", "Type./Sp. figured.", andasmalllabelwithamalesymbol; andaround whitelabelwith a red border printed with "Type/H.T". Male. July-October, size = 10.9 (10.4-11.1, N = 13). Dorsum blackish-brown (Sepia, color 119); marginal row ofblackdotsoftenwith faintwhitepointsproximally especially towards forewing apex; forewing with four quadrate subapical white spots, continued submarginally as irregular row of small and poorly defined white spots; large quadrate whitebaratendofdiscalcellwithsmaller, lessdistinctbarbelowitincellCuA,;postbasal white bar in discal cell and similar onebelow it in cell CuA2; subapical, submarginal and cell-end white marks indistinctly outlined distally and proximally with black; similarly, postbasalbaroutlined distally andbarbelow cell-end bar outlined proximally. Hindwing with markings as forewing except entire submarginal row of approximately equal-sized 136 Journal of the Lepidopterists' Society Figs. 1-8. Apodemia hepburni subspecies (dorsal surface on left, ventral surface on righWt of each figure). 1-4, A. h. hepburni (all MEXICO: Sonora)W—(1) male, Rt. 16, 8.5 mi RioYaqui, 26 Aug. 1984, leg. D. Mullins, (2) female, 36mi Moctezuma, 3 Aug. 1984, leg. D. Mullins, (3) male, Hwy. 16, 17 mi E Tecoripa, 15 March 1984, leg. J. P. Broc—k, (4) female, same data as fig. 3; 5-8, A. h. remota (all MEXICO: Baja California Sur) (5) holotype, data in text, (6) allotype, data in text, (7) male, Mulege, Vz mi E, 23 March 1974, leg. G. S. Forbes, (8) female, 3 mi S Loreto, leg. Faulkner, Brown. spots;slightreddish-brownareaatmiddleofcostalmargin,sometimesextendingmedially into discal cell; fringes of both wings white, lightly checked with black at vein tips. Ventral surface dull reddish-brown; forewing near Pratt's Rufous proximally grading to Robin Rufous distally; hindwing Pratt's Rufous throughout; hindwing and base of forewing overscaled with white (often heavily); white marks as on dorsum but larger, especially submarginally; submarginal row of hindwing forming continuous, irregular Volume 45, Number 2 137 band; white spots outlined with black as on dorsum but more prominent against paler ground color; marginal black dots with indistinct white smudges proximally; hindwing anal margin whitish. March, size = 11.7 (10.9-12.4, N = 7). Dorsum paler (near Hair Brown), forewing subapical and cell-end white marks prominent; posterior spots faint to nearly obsolete, position indicated by their black outlines; hindwing white marks similarly faint, most persistentinsubmarginalseries;marginalblackspotssmall,nearlydisappearinganteriorly on forewing; hindwing flush of reddish-brown tending more prominent. Ventral surface paler (nearSalmonColor, color 106) proximally, grayerdistally;blackmarkingsobsolete; white marks less contrasting. Female. August-October, size = 11.1 (10.3-11.8, N = 14). Wings more rounded than male; groundcolor asmale; white marksasmale; forewing with indistinct Pratt's Rufous margin; marginalblackspotswith proximal indistinct points ofwhite apically; hindwing with anterior Va and margin Pratt's Rufous. Ventralsurfacesimilartomalebutslightlypaler; ground colornearCinnamon Rufous throughout, less overscaled with white except between marginal black dots and submar- ginal white spots giving appearance of whitish band. March, size = 12.2 (11.9-12.4, N = 2). As male, anterior Vz of hindwing with reddish- brown flush, forewing with more restricted marginal reddish-brown than later in year. Distribution andphenology. The distribution of this taxon apparently is incompletely known. Nearly all records that I know of are from Sonora, Mexico (Fig. 9) where it may be locally common. It was described from western Chihuahua, Mexico; there are old records from southern Arizona (various museums), additional, more recent, Arizona re- cords from northeast of Douglas, Cochise County (fide R. Bailowitz) and at Sycamore Canyon, Santa Cruz County (Langston 1991), and it was taken in the Chisos Mountains, Brewster County, Texas (Carnegie Museum). The flight period in Sonora is from late FebruarytolateMarchandlateJulytolateOctoberreflectingatleasttwobroods. Arizona records are for May and June (one in August) and the Texas record is for July. Discussion. This taxon isat leastbivoltine in spring and again in autumn andstrongly biphenic. The springbrood is pale with reduced markings; the fall phenotype is smaller, darker and the ventral surface is relatively heavily overscaled with white and appears mottled. Apodemia hepburni remota, new subspecies (Figs. 5-8) Male. August-November, size = 11.6 (10.2-12.6, N = 25). Dorsum darkbrown (Sepia, color219);submarginalrowofblackspotsonbothwingsasonA. h. hepburnibutwithout associatedwhitepoints;whitemarksandblackoutlinesasonA./i. hepburnibutnarrower; hindwing usually with medial flush of reddish-brown anteriorly, extending into discal cell. Ventralsurfacenearlyuniform Pratt'sRufouswithlittlewhiteoverscaling; whitespots asondorsum,theirblackoutlinesthin;blackmarginaldotssmall,oftenobsoleteanteriorly on forewing, and with associated white very vague or absent. February-April, size=12.3(11.9-13.4, N=7). Dorsalgroundcolorpaler(HairBrown), posteriorwhitemarksonforewingandbasalmarksonhindwingfainttoobsolete;marginal black spots very small (maybe absent on forewing); reddish-brown on hindwing present or absent. Ventralgroundcolorpale(nearOrange-Rufous); verylittlewhitishoverscalingbasally, white marks very thin withblack outlines faint toobsolete; marginal black spots reduced to points or entirely absent. Female. August-November, size = 12.4 (11.0-14.1, N = 30). Wings more rounded than male; hindwing with relatively heavy flush of reddish-brown anteriorly; margins with smudges of similar color in each cell; white markings thin as on male. Ventral surface paler than male (near Orange-Rufous); markings as on male. April, size = 12.6 (12.3-12.9, N = 2). Dorsum similar to later in year but tending 138 Journal of the Lepidopterists' Society towardsmorereddish-brown onhindwing. Ventral surfacevery pale (near Flesh Ocher), markings as on maleM.— Types. Holotype MEXICO: BajaCaliforniaSur—;ArroyoSanBartolo, 28Aug. 1982, leg. [J. W.] Brown and [D. K.] Faulkner. Allotype F same data as holotype. Paratypes (60M, 50F, all MEXICO: Baja California Sur)—same data as holotype (1M, 10F); same location as holotype, 28 Nov. 1980, leg. J. Brown (IF); San Bartolo, 3 Oct. 1981, leg. F. Andrews & D. Faulkner (2F); 30 Nov.-l Dec. 1979, leg. Brown & Faulkner (2M, 3F); A. San Bartolo, 2 Nov. 1961, leg. Cary-Carnegie Expedition [=CCE] (IF); 3 Nov. 1961, leg. CCE (3M, 2F); 12 Nov. 1961, leg. CCE (IF); San Bartolo microwave tower, 28 Nov. 219m80i,SlWeg.CJa.dBuarnoow,n2(61MA)u;g.31m9i82S,EleSga.nFaBuarltkonleor, &15BMraorwcnh(719M7,4,IFl)e;g.7Gk.mS.SFCoardbueasno(,1M2);6 Aug. 1982, leg. Faulkner and Brown (1M, IF); Caduano, 25 Nov. 1961, leg. CCE (1M); Ro. Palmarito 4 Nov. 1961, leg. CCE (2F); 5 Nov. 1961, leg. CCE (1M), 30 Nov. 1961, leg. CCE(IF);31 Nov. 1961, leg. CCE(1M, IF);RanchoSanBernardodeSierraLaguna, 14 Nov. 1961, leg. CCE (1M, IF); 17 Nov. 1961, leg. CCE (1M); Bahia de Palmas, 20 Nov. 1961, leg. CCE (1M, 2F); 33 mi N Todos Santos, 4 Oct. 1981, leg. D. Faulkner & F. Andrews (1M); 28 km N Todos Santos, 29 Nov. 1980, leg. Brown & Brown (IF); 31 km N Todos Santos, 29 Nov. 1980, leg. J. Brown (IWF); 4 mi E La Barrera (nr. Todos Santos), 21 March 1974, leg. R. Holland (W1M); 8 mi La Paz, 30 Oct. 1946, leg. E. Y. Dawson (1M); Ramel de Naranjas, 6 mi Hwy. 1 nr. Santa Anita, 11 Oct. 1983, leg. Andrews & Faulkner (2M); San Antonio microwave, 13 Oct. 1983, leg. D. Faulkner & F. Andrews (3F); Boca de la Sierra, 13 Nov. 1961, leg. CCE (4M); 17 Nov. 1961, leg. CCE (4M, IF); 22 Nov. 1961, leg. CCE (1M, IF); 24 Nov. 1961, leg. CCE (2M, 2F); 28 Nov. 1961,leg.CCE(3M,2F);PuertoChileno,22Nov. 1961,leg.CCE(2M,IF);Guaycura Hotel, La Paz, 29 Nov. 1961, leg. CCE (1M); 15 mi S La Paz, 1 Nov. 1946, leg. E. Y. Dawson (16M, 2F); 2 mi S Buena Vista, 30 Nov. 1979, leg. Brown & Faulkner (IF); 5 kmSRBuenavista, 25Oct. 1961, leg. CCE (2F);Santiago, 6Nov. 1946, leg. E. Y. Dawson (2F); 3 mi S Santiago, 25 Oct., leg. ? (IF); San Jose del Cabo, 23 Nov. 1961, leg. CCE (IF). Deposition of types. The holotype, allotype, and 58 paratypes are deposited at the NaturalHistoryMuseum,SanDiego,California;49paratypesareattheCarnegieMuseum of Natural History, Pittsburgh, Pennsylvania; one paratype is in the private collection of G. S. Forbes, Las Cruces, New Mexico, and two paratypes are retained by the author. Type locality. MEXICO: Baja California Sur; Arroyo San Bartolo. San Bartolo is on Mexico Highway 1 between La Paz and San Jose del Cabo. Distribution and phenology. Apodemia h. remota occurs only in Baja California Sur from Mulege south to the southern tip (Fig. 9). Specimens have been taken from mid February to mid April and from late August to early December, about 70% (of 117 examined) in November. Etymology. Thissubspeciesisnamedafteritsisolateddistribution atthetipoftheBaja California peninsula. Diagnosis and discussion. Apodemia h. remota is seasonally biphenic in size, color, and pattern; thisissimilarinformbutsomewhatlesspronounced thanfornominotypical A. hepburni. Apodemia h. rmemmota differs in several respects frommmA. h. hepburni. It is a larger insect (averaging 0.5 larger in the first brood and 1 in the second), both theventralanddorsalgroundcolorsarepaler, thewhitemarkingsandtheirblackoutlines are thinner (especially on the ventral surface where they are about twice asbroad on A. h. hepburni), the black marginal spots are smaller, and there is considerably less whitish overscaling on the ventral surface (see Figs. 1-8). Overall, the ventral surface of A. h. remota appears uniformly colored; that of A. h. hepburni has a mottled appearance, especially on late season specimens. Apodemia h. remota tends to have more red-brown medially on the dorsal hindwing (this sometimes extends distally to the outer margin on females) and along the margins on both wings than A. h. hepburni and rarely has white associated with the black marginal spots. Itiscuriousthatmostreferencesdonotmention theoccurrenceofA. hepburniinBaja California Sur. Hoffmann (1976) acknowledged theoccurrenceof thespeciesonlyonthe mainland. He, Rindge (1948), andHolland (1972) reportedonlyA.palmerii(=Apodemia Volume 45, Number 2 139 ARIZONA >v"s*,>^oo X T" \TEXAS SONORA o o CHIHUAHUA CD °oco BAJA CALIFORNIA SUR Fig. 9. Known distribution of Apodemia hepburni. Open circles = A. h. hepburni, closed circles = A. h. remota, "x" = specimens not seen. murphyi Austin) of this group in Baja California. Of other general references, Howe (1975) reported its occurrence in Baja California but Scott (1986) did not. Discussion Both taxa of A. hepburni are relatively rare in collections although the species appears to be common locally. The dates of collection seem to reflect two well defined and disjunct flight periods in both Sonora and Baja California Sur, Mexico. It is unknown how much of the known phenology reflects collector phenology. In Baja California, however, there are records for A. murphyi from every month, suggesting that the absence of A. hepburni records in some months is real. The few records for Arizona and Texas bridge the phenological gap in Mexico, perhaps reflecting a single brood at the northern extreme of the dis- tribution. I have found no information on the early stages or larval food plant of A. hepburni. Seasonal variation of both A. hepburni taxa is parallel. Individuals from early in the year (spring) are large and pale with small white markings;thosefrom laterintheyear (latesummer-fall) areofasmaller, 140 Journal of the Lepidopterists' Society Table 1. Characteristics of species of the Apodemia palmerii complex. Character A.palmerii A.murphyi A.hepburni FW shape not produced produced produced FW apex rounded subfalcate subfalcate Dorsal margin prominent series of prominent series of no or inconspicuous white spots white spots white spots Dorsal macula- prominent prominent vague, esp. posteri- tion orly on forewing VHW postmedi- medium width very broad narrow to medium an band width Distribution SW US to central southern Baja Cali- southern Baja Cali- Mexico fornia, Mexico fornia and west- ern Mexico darker phenotype with more extensive white spots. Seasonal biphenism wasnoted in sizeand color ofA. p. palmeriiand in colorforA. murphyi although none was detected for two other A. palmerii taxa (Austin 1987). Apodemia hepburni occurs sympatrically and synchronically with A. palmerii or A. murphyi; the two have been taken together at several locations in Sonora and Baja California. The known distribution and flight period of A. palmerii and A. hepburni are practically identical in Sonora. In Baja California, A. hepburni has a shorter flight period and does not extend as far north as A. murphyi. Apodemia hepburni is sometimes confused (in collections) with A. palmerii. Apodemia palmerii always has a prominent row of white spots just proximal to the marginal black dots on both wings; these are absent or represented at most by inconspicuous white points on A. hepburni. The white marks of the submarginal series on the ventral surface are usually conspicuously outlined with black on both sides on late season A. hepburni; this black outline is usually absent distally on spring A. hepburni and all A. palmerii (some A. palmerii may have a darker shade here but never a well-defined line of black). Additionally, the forewings of both sexes of A. hepburni are more produced apically than those of A. palmerii. Apodemia h. remota likewise resembles A. murphyi. The wing shape of the two is similarly produced. Apodemia hepburni, however, has smaller dorsal spots, no reddish-brown basally on the forewing (occasional female A. hepburni have a hint of this but it is never prominent as on some A. murphyi), lacks the marginal white spots, and has a very narrow white postmedian band on the ventral hindwing (very broad and prominent onA. murphyi). These characters are summarized in Table 1. The malegenitaliaof the twosubspeciesofA. hepburni are identical; they are also very similar to those of A. palmerii. The processes of the Volume 45, Number 2 141 valvae diverge at a greater angle on A. hepburni than on A. palmerii and the lower process is slightly shorter and twisted outward. The vinculum ofA. hepburniappears more upright compared tothesloping aspectofA. palmerii. The malegenitaliaofA. murphyi are more robust than those of either A. hepburni or A. palmerii. Acknowledgments Ithank variouscuratorsandothersforloansofspecimens: L. D. andJ. Y. Miller(Allyn Museum of Entomology), F. H. Rindge (American Museum of Natural History), D. K. Faulkner(NaturalHistoryMuseum, SanDiego),J. P. Donahue(NaturalHistory Museum of Los Angeles County), R. K. Robbins (National Museum of Natural History), J. E. Rawlins (Carnegie Museum of Natural History), A. Powell (California Insect Survey), and T. W. Davies (California Academy of SciencJe.s). I am grateful for specimen loans and data from the private collections of J. Brock, G. S. Forbes, R. W. Holland, R. O. Kendall, D. Mullins, and M. J. Smith. I also thank P. R. Ackery of the Natural History Museum, London, for sending photographs of the type of A. hepburni. I thank Scott Klette of the Nevada State Museum for the photographs in the text. Literature Cited Austin, G. T. 1987. Apodemia palmerii (Lycaenidae: Riodininae): Misapplication of names, two new subspecies and a new allied species. Res. Lepid. 26:125-140. J. Godman, F. D. & O. Salvin. 1879-1901 [18861 Biologia Centrali-Americana. Lepi- doptera: Rhopalocera. Vol. 1. London, xlv + 487 pp. Hoffmann, C. C. 1976. Catalogo sistematico y zoogeografico de los lepidopteros mex- icanos. Soc. Mex. Lepid., publ. esp. 1. 214 pp. Holland, R. 1972. Butterflies of middle and southern Baja California. Res. Lepid. J. 11:147-160. Howe, W. H. 1975. The butterflies of North America. Doubleday, Garden City, New York, xiii + 633 pp. Langston, R. L. (coordinator). 1991. 1990 season summary. Southwest: Arizona, Cal- ifornia, Nevada. News Lepid. Soc. 1991:16-19. Miller,L. D. & F.M. Brown. 1981. Acatalogue/checklistofthebutterfliesofAmerica north of Mexico. Lepid. Soc. Mem., No. 2. vii + 280 pp. Rindge, F. H. 1948. Contributions toward a knowledge of the insect fauna of Lower California. No. 8, Lepidoptera: Rhopalocera. Proc. Calif. Acad. Sci. 24:289-312. Scott, J. A. 1986. The butterflies of North America: A natural history and field guide. Stanford Univ. Press, Stanford, California, xiii + 583 pp. Smithe, F. B. 1975. Naturalist's color guide. Amer. Mus. Nat. Hist., New York, New York. 4 pp. + 8 color charts. 1981. Naturalist's color guide part III. Amer. Mus. Nat. Hist., New York, New York. 37 pp. + 9 color charts. Receivedfor publication 8 March 1990; revised and accepted 13 June 1991.